Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 November 2025 | 17(11): 27863–27873
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.9820.17.11.27863-27873
#9820 | Received 07 April 2025 | Final received 31 October 2025 | Finally
accepted 17 November 2025
Phylogenetic confirmation of
generic allocation and specific distinction of Mawphlang Golden-cheeked Frog Odorrana
mawphlangensis (Pillai & Chanda, 1977) (Amphibia: Anura: Ranidae) and
its updated distribution records
Angshuman Das Tariang 1,
Mathipi Vabeiryureilai 2, Fanai Malsawmdawngliana 3 & Hmar Tlawmte Lalremsanga
4
1,2,3,4 Developmental Biology and
Herpetology Laboratory, Department of Zoology, Mizoram University, Aizawl,
Mizoram 796004, India.
1 angshumanjay29@gmail.com, 2 mvabeiryureilai@gmail.com,
3 sawmattr.yx@gmail.com, 4 htlrsa@yahoo.co.in
(corresponding author)
Abstract: We report on the phylogenetic position (16S rRNA) of the Mawphlang
Golden-cheeked Frog Odorrana mawphlangensis collected from
Meghalaya (topotypical) and Mizoram, northeastern India. Morphologically, our
new specimens agreed with the original description and subsequent redescription
of O. mawphlangensis, thus ratifying the generic placement of O.
mawphlangensis and its specific distinction from O. grahami,
to which it was hypothesized to be a synonym owing to morphological
similarities. The observed interspecific genetic distances within the genus Odorrana
from our study ranged from 0.00% (O. mawphlangensis) to 11% (O.
fengkaiensis). The uncorrected K2p-distance of 16S rRNA among Odorrana
species revealed O. mawphlangensis is closest to its sister
species O. jingdongensis, with a genetic distance of 3.7%. This
study identifies and confirms the collected samples from Mizoram as O.
mawphlangensis and also presents an updated distribution record from
northeastern Indian state of Mizoram apart from its type locality in Meghalaya.
Keywords: Distribution records, frog,
holotype, type locality, mitochondrial DNA, Mizoram, molecular phylogeny,
morphology, northeastern India, taxonomy.
Editor: S.R. Ganesh, Kalinga
Foundation, Agumbe, India. Date of publication: 26 November 2025 (online & print)
Citation:
Tariang, A.D., M. Vabeiryureilai, F. Malsawmdawngliana & H.T. Lalremsanga (2025). Phylogenetic
confirmation of generic allocation and specific distinction of Mawphlang
Golden-cheeked Frog Odorrana mawphlangensis (Pillai & Chanda, 1977)
(Amphibia: Anura: Ranidae) and its updated distribution records. Journal of Threatened Taxa 17(11): 27863–27873. https://doi.org/10.11609/jott.9820.17.11.27863-27873
Copyright: © Tariang et al. 2025. Creative Commons Attribution 4.0 International License.
JoTT allows unrestricted use, reproduction, and distribution of this article in
any medium by providing adequate credit to the author(s) and the source of
publication.
Funding:
(i) MZU UGC Non-NET Fellowship, 2-5/MZU(Acad)/22(Ph.D)/70 to Angshuman Das
Tariang. (ii) DST-SERB, New Delhi (EEQ/2023/ 000877) to Dr. Mathipi
Vabeiryureilai. (iii) Fellowship and Scholarship for Higher Education of ST
Students under the Ministry of Tribal Affairs, Government of India, New Delhi
File no. 202122-NFST-MIZ-00046 to Fanai Malsawmdawngliana. (iv) DST-SERB, New
Delhi (CRG /2023/000805), the Indian Council of Medical Research (ICMR Nos.
58/6/2/SB-CFP/AKM/2023/NCD-II and 58/6/3/SB-CFP/AKM/2023/NCD-II) to Prof. Hmar
Tlawmte Lalremsanga.
Competing interests:
The authors declare no competing interests.
Author
details: Angshuman Das Tariang is a Ph.D. research scholar in the
Department of Zoology, Mizoram University working under the supervision of
Prof. H.T. Lalremsanga. He specializes in the field of developmental biology, herpetology,
taxonomy and phylogenetics. He is currently working on anurans of Meghalaya. Dr. Mathipi Vabeiryureilai is an
assistant professor in the Department of Zoology, Mizoram University. He
specializes in Cancer Research, Zoology and Physiology. He is currently working
on DNA barcoding of amphibians and reptiles from Mizoram, India. Fanai Malsawmdawngliana is a Ph.D.
research scholar in the Department of Zoology, Mizoram University working under
the supervision of Prof. H.T. Lalremsanga. He specializes in the field of
systematics, molecular phylogenetics and biogeography.
Prof.
Hmar Tlawmte Lalremsanga
is the head of the Department of Zoology, currently teaching PG students and
guiding 14 PhD scholars and looking after 6-major projects. He specializes in
diversity, morphology and molecular biology of insects, amphibians and
reptiles. He has worked extensively on ecological survey on breeding and
development in relation to environmental factors. He excels in systematics
(taxonomy), ecology and developmental biology.
Author contributions: ADT
conceived and led the study, undertaking the collection of specimens,
conducting detailed morphological examinations, taking photographs of the
samples and drafting the complete manuscript. MV was responsible for executing
the molecular analyses and contributed to the interpretation of genetic data.
FM developed the geographical distribution map and
assisted in visual representation of the findings. HTL provided critical
scholarly guidance throughout the study, offered substantive inputs to
strengthen the manuscript, and oversaw the overall research process. All
authors contributed to refining the final version of the manuscript and
approved its submission.
Acknowledgements: Collecting
permit issued by the Environment, Forests and Climate Change Department,
Mizoram, Government of Mizoram, India (No.A.33011/2/99-CWLW/225) and Office of
The Principal Chief Conservator of Forests, Wildlife & Biodiversity and
Chief Wildlife Warden, Meghalaya, Government of Meghalaya, India
(FWC/Research/118/3460-65).
INTRODUCTION
Odorrana
mawphlangensis (Pillai & Chanda, 1977) commonly known as the Mawphlang
Golden-cheeked Frog (Dinesh et al. 2023), is a large sized frog species
belonging to the family Ranidae, first described by Pillai & Chanda (1977),
from Mawphlang Sacred Forest in Khasi Hills, Meghalaya, India, on the basis of
an adult female bearing museum number, ZSI A 6979 (ex ZSI/ERS 803). In
subsequent observations, this species was reported from other northeastern Indian
states of Nagaland, Arunachal Pradesh, Manipur, and Mizoram as well as from
West Bengal (Frost 2025). The description of the species was based solely on
the holotype which led to it being misleadingly placed as a close relative of
several species, namely Limnonectes doriae (Boulenger, 1887) and Limnonectes
modestus (Boulenger, 1882). This misplacement was followed in the
subsequent reports by Tiwari (1981) and Dubois (1987) with Ohler & Dubois
(1999) transferring this species to the genus Limnonectes (Elachyglossa)
(Anderson, 1916), including the originally proposed “possible relative”
of L. modestus. Ao et al. (2003) indicated its close resemblance to Rana
(Odorrana) andersoni. However, there was not enough
explanation to support this inference. Mahony (2008), encountered
similar-looking ranid frogs at the type locality Mawphlang Sacred Forest that
not only resembled Odorrana but also produced secretions having a
distinct odour. Against the back drop of these sightings, Mahony (2008)
redescribed the holotype (ZSI/K) which re-allocated it from Limnonectes, to
Odorrana based on morphological and geographical grounds.
The Ranid
genus Odorrana comprises of 69 recognized species (Frost 2025), widely
distributed in montane streams and rivers in the subtropical and tropical
regions of eastern and southeastern Asia (Fei et al. 2012; AmphibiaChina 2021;
Frost 2025). High-gradient streams, often located in mountain environments, are
typical habitats of the Odorrana species, although O. mutschmanni (Pham
et al. 2016), O. wuchuanensis (Xu 1983), and O. lipuensis (Mo et
al. 2015) have been found to otherwise inhabit karstic limestone caves, with an
elevation range of 447–728 m (Fei et al. 2012; Pham et al. 2016; Liu et
al.2021; Frost 2025). The distributional range of the genus comprises of the
Ryukyu Archipelago (Japan), southern China, northeastern India, and the
Thai-Malay Peninsula, and further extending southwards to the two large southeastern
Asian islands, Sumatra and Borneo (Frost 2025). Considering the peculiar
ecological niche where this species resides, there could possibly be inter- or
intra-specific divergence through geographical isolation, especially given the
species complex with a wide geographical range covering varied mountain ranges
and variable habitats (Wang et al. 2015). The phylogeny and diversity of Odorrana
and the systematic status of taxa within the genus have been a matter of
constant debate by taxonomists (Frost et al. 2006; Che et al. 2007; Fei et al.
2009; Kurabayashi et al. 2010; Chen et al. 2013; Li et al. 2015).
In spite of
treating Odorrana as a subgenus of Rana (Dubois 1992) and
expanding the genus Huia to include both Odorrana and Rana
(Eburana) (Frost et al. 2006), the monophyly of Odorrana was
finally supported by analysing mtDNA data and nuclear data separately as well
as subsequent analyses of the combinations of mtDNA and nuclear data. Thus, the
controversies revolving around the systematic status of Odorrana to be
considered as a subgenus of Rana (Eburana), as well as being
included with Rana as subgenus of Huia were subsequently
abandoned (Matsui et al. 2005; Stuart 2008; Pyron & Wiens 2011). Recent
phylogenetic studies have confirmed that Odorrana is monophyletic and
consists of at least seven clades (Chen et al. 2013). In this study, we
successfully conducted a detailed phylogenetic analysis and assessment of O.
mawphlangensis based on 16S rRNA in addition to studying the morphological
parameters of the collected samples to resolve the confusion revolving around
its taxonomic position and identity.
MATERIALS
AND METHODS
Survey and
morphometric analysis
Visual
encounter surveys (VES) were conducted in 2022 and 2023 (August to September)
and a total of 12 specimens were collected during the field surveys from the
Indian states of Mizoram and Meghalaya. Ten specimens were collected from
Mizoram and two specimens were collected from Meghalaya (Figure 1). Out of the
10 specimens obtained from Mizoram, eight specimens were found from Murlen
National Park (23.673o N, 93.293o E; 1,050 m) and two
specimens were obtained from Hmuifang Community Reserve Forest (23.355o
N, 92.753o E; 1,458 m). The other two specimens of Meghalaya were
collected from Malki Forest in East Khasi Hills District (25.562o N,
91.893o E; 1,549.5 m). They were found in their natural habitats,
i.e., on rocks near the riverbeds and in seasonal intermittent stream below
hilly grass-covered terrain (Frost 2025). The specimens were deposited,
catalogued, and fixed in 70% ethanol solution in the Departmental Museum of
Zoology, Mizoram University (MZMU). Individuals were identified using the
literature of Kiyasetuo & Khare (1986) and Chanda (2002). Morphometric
measurements (Table 1) follow Sengupta et al. (2010). Measurements to the nearest 0.1 mm were taken
using a dial calliper (MitutoyoTM 505-507) (Table 1). Photographic vouchers
were submit-ted to the Natural History Museum of Mizoram, Mizoram University,
Aizawl, Mizoram, India (NHMM/G/2–4).
DNA
extraction, PCR amplification and sequencing
We extracted
genomic DNA from the liver tissues of Odorrana using
QIAamp DNA Mini Kit (Qiagen Cat. No. 51306) following the standard protocol
provided by the manufacturer. DNA integrity was analysed on 0.8% (w/v) agarose
gel containing ethidium bromide. Polymerase chain reaction (PCR) was prepared
for 20 μL reaction mixture containing 1X amplification buffer, 2.5 mM MgCl2,
0.25 mM dNTPs, 0.2 pM each forward and reverse primer, 1 μL genomic DNA, and 1U
Taq DNA polymerase with a pair of partial 16S rRNA primers: forward (L02510-
CGC CTG TTT ATC AAA AAC AT) (Palumbi 1996) and reverse (H03063- CTC CGG TTT GAA
CTC AGA TC) (Rassmann et al. 1997). The PCR thermal regime for amplification
was 5 min at 95°C for initial denaturation, followed by 35 cycles of 1 min at
95°C for denaturation, 30s for annealing at 50.3°C, elongation for 1 min at
72°C, and a final elongation for 5 min at 72°C. Amplicons were observed through
gel electrophoresis using a 1.5% agarose gel containing ethidium bromide.
Samples were sequenced using Sanger’s dideoxy method and sequencing was carried
out for both forward and reverse directions (Barcode Bioscience, Bangalore,
India). The newly generated partial 16S rRNA sequences were deposited in the
GenBank repository to obtain the accession number.
Phylogenetic
analyses
The
phylogenetic relationships among the genus Odorrana were assessed based
on the 16S rRNA partial gene sequence. For the analysis of our dataset of 16S
rRNA, we included three newly generated O. mawphlangensis sequences from
Mizoram (MZMU 138, MZMU 139, & MZMU 2267) along with the sequence of the
holotype samples collected from Meghalaya (MZMU 3020 & MZMU 3021); 21
sequences were retrieved from NCBI database, and one sequence of Amolops
indoburmanensis (MT790757) sample was used for this study as an out-group.
The sequences were aligned by using Muscle algorithm in Molecular Evolutionary
Genetics Analysis 7 (MEGA 7) (Kumar et al. 2016), the mean uncorrected genetic
distances (Kimura 2 parameter, K2P) (Kimura 1980) were calculated in MEGA 7
(Kumar et al. 2016). The final aligned dataset contains 535 positions of 16S
rRNA gene sequences. The best-fitting models of DNA evolution was performed
according to the modeltest - NG (Darriba et al. 2020), and selected
GTR+G model suggested by models with the lowest Bayesian information criterion
(BIC) and Akaike information criterion (AIC) scores. The phylogenetic tree was
inferred using Bayesian inference (BI) and maximum likelihood (ML) approaches.
ML analysis was performed in raxmlGUI-2 with one-thousand bootstrap replicates
(Silvestro & Michalak 2012). The BI phylogenetic tree was reconstructed in
MrBayes 3.2.5 using GTR+G model. The Markov Chain Monte Carlo (MCMC) (one cold
and three hot chains) was run for 10,000,000 generations by sampling every 100
generations and set the burn-in to 25%. The analysis was terminated when the
standard deviation of split frequencies was less than 0.001. The percentage of
trees in which the associated taxa clustered together is shown next to the
branches (Ronquist & Huelsenbeck 2003). The generated phylogenetic tree was
further illustrated using Figtree v1.4.4 software (Rambaut 2018).
Genetic
distances
The
resulting sequences were deposited in GenBank depository (OP979109, OP979112,
MZ229896, MT814039, & MT814038). The GenBank accession numbers and other
details of the species involved in the phylogenetic Bayesian inference tree
have been presented in Figure 2. The uncorrected pairwise divergences in the
16S rRNA gene fragment among the members of the Odorrana sp. were
summarised in Table 3. The observed interspecific genetic distances within the
genus Odorrana from our study ranged from 0.00% (O. mawphlangensis)
to 11.00% (O. fengkaiensis_KT31538). Moreover, an intraspecific genetic
distance of O. mawphlangensis was detected in the range of 0.00–0.011
between the specimens from Mizoram and type locality, i.e., Meghalaya.
RESULTS
a)
Morphology (also see Mahony 2008)
Odorrana
mawphlangensis has some distinguishing morphological characters which are exclusive to
this species, such as: the head length > head width, snout length > eye
diameter, inter-orbital distance < internasal distance, forelimb length <
hand length, colouration of the body, which is dorsally
primarily green anteriorly, with large randomly spaced brown spots, lip-stripe
yellow to bronze/brown in adult. The body colouration of the collected samples
was green at the anterior part of the dorsum with large brownish spots randomly
spaced and the posterior part of the dorsum was brownish in colour. In all the
collected specimens, it was observed that the head is longer than its width
(i.e. HL > HW) with the HL/HW ratio of 22.12 : 17.51 and the snout length is
longer than the eye diameter (i.e., SL > ED) with the SL/ED ratio of 6.57 :
5.58. Moreover, it was observed that the inter-orbital distance is less than
the internasal distance and the length of the forelimb is shorter than the
hindlimb length. All these observed morphological parameters indicated that the
collected samples were O. mawphlangensis. The tympanum diameter (TYD)
constituted 60.24% of the eye length (EL). Image 2 depicts a preserved sample
of O. mawphlangensis (MZMU 138) collected from Mizoram.
From our
collected specimens of O. mawphlangensis, morphometric analyses resulted
in the following comparisons (vs. Chanda 1994; Ao et al. 2003; Mahony 2008):
Snout-vent
length (SVL) was in the range of 31.8–94.4 mm. The largest male among the
collected specimens had SVL of 80.0 mm. Chanda (1994) provided SVL in the range
of 60.0–90.0 mm. Ao et al. (2003) reported SVL of 80.0 mm in males and a range
of 96–106 mm for females while Mahony (2008) redescribed the holotype from
Mawphlang Sacred Forest by reporting an adult female with SVL of 84.3 mm.
Head width
(HW) ranged from 11.2–31.7 mm and Head length (HL) ranged from 14.2–41.9 mm. As
per Mahony (2008), HW was reported as 31.0 mm and HL as 34.4 mm.
The range of
eye diameter (ED) or eye length (EL) was 3.7–11.2 mm and Mahony (2008) reported
ED of 10.1 mm.
Internasal
distance (IN) was in the range of 1.7–9.9 mm whereas Mahony (2008) reported it
at 10.8 mm.
Eye-nostril
distance (EN) was in the range of 1.9–6.2 mm against 7.1 mm as per Mahony
(2008).
The greatest
tympanum diameter (TYD) was in the range of 1.9–5.4 mm against 5.3 mm by Mahony
(2008).
The mean
inter-orbital distance (IUE) was found to be 3.7 mm which is shorter than the
mean internasal space (IN), i.e., 4.4 mm (i.e., IUE < IN).
The forelimb
length (FLL) ranged 5.2–19.7 mm and was shorter in length than the hand length
(HAL) which was in the range of 8.0–26.0 mm (i.e., FLL < HAL).
Tibia length
(TL) was observed to be longer than the femur length (FL) (i.e., TL > FL),
ranging 16.0–49.7 mm and 14.8–40.1 mm, respectively. Mahony (2008) had reported
TL of 51.6 mm and FL of 47.5 mm.
b) Field observations and comparisons
The largest
specimen was a female, catalogued as MZMU 138. From our collected specimens of O.
mawphlangensis, morphometric analyses have revealed that the largest
specimen was obtained from Hmuifang Community Reserve Forest with SVL of 94.4
mm (an adult female) and the smallest specimen belonged to Murlen National Park
with SVL of 31.8 mm (a juvenile). A number of adult specimens of O.
mawphlangensis that were encountered at Malki Reserve Forest and Hmuifang
Community Reserve Forest (Image 1A,B). Amplecting behaviour between two adults
was also observed (Image 1A).
c) Phylogenetic Relationships
In
phylogenetic reconstructions (Figure 2) using both BI and ML analyses, the
trees revealed identical topologies that support the relationships of the genus
Odorrana. All 17 sequences of Odorrana sp. were clustered and
formed distinct monophyletic clade from an out-group, Amolops
indoburmanensis (Figure 2). The samples used for phylogenetic
analysis have been mentioned in Table 2. From our analysis, Odorrana
mawphlangensis nested different from other species with well supported
bootstrap values (Figure 2; Table 3).
d) Conservation
Status
With the recent amendment of the Wildlife Protection Act in 2022, 37
species of amphibians have been protected under Schedule-I and Schedule-II
categories. Odorrana mawphlangensis has been included in Schedule-II
category. This species is currently listed as ‘Data Deficient’ and needs
further research and assessment (IUCN 2023).
DISCUSSIONS
There were
numerous brief accounts and reports on the occurrence of this species in the
past (Chanda 1990; Sarkar et al. 1992; Dutta 1997; Chanda 2002; Ao et al. 2003;
Devi & Shamungou 2006; Sarkar & Ray 2006; Das & Dutta 2007; Ahmed
et al. 2009; Matthew & Sen 2010; Lalremsanga 2017; Siammawii et al.
2021). However, these studies were reports solely based on the external
morphological characters and a detailed phylogenetic analysis was not performed
in any of the studies. Thus, there was confusion on its identity and a detailed
confirmation was awaited.
Mahony
(2008) redescribed the species based on re-examination of its holotype,
illustrated the same and transferred it to the genus Odorrana. Mahony
(2008) also suggested that it might be conspecific with O. grahami, based
on some morphological similarities. He stated that further studies on the
phylogenetic position and distinction are required so as to confirm the
monophyly as well as establishing the taxonomic identity of the species
involved.
In the
present study, based on 16s rRNA data, we ratify the generic allocation of Odorrana
mawphlangensis (see Mahony 2008) and establish that its closest sister
taxon is the Yunnanese species O. jingdongensis (Fei et al. 2001) with a
genetic distance of 3.7%. The hypothesis that O. mawphlangensis might be
a junior synonym of O. grahami (Mahony 2008) is not supported from this
study. It is noteworthy that O. mawphlangensis forms a distinct clade
separate from sister taxa O. grahami and O. jingdongensis
(Boulenger 1917; Fei et al. 2001).
Table 1. Morphometric
measurements (in mm) of the specimens of Odorrana mawphlangensis
collected from Mizoram (Murlen National Park and Hmuifang Community Reserve
Forest) and Meghalaya (Malki Reserve Forest).
|
Locality |
Murlen NP |
Hmuifang RF |
Malki RF |
|||||||||
|
Museum number |
MZMU 1836 |
MZMU 1995A |
MZMU 2129 |
MZMU 2856 |
MZMU 2858 |
MZMU 2859 |
MZMU 2861 |
MZMU 2866 |
MZMU 138 |
MZMU 139 |
MZMU 3020 |
MZMU 3021 |
|
SVL |
57.3 |
54.6 |
43.2 |
38.8 |
36.3 |
31.8 |
31.9 |
32.8 |
80.0 |
94.4 |
44.3 |
46.1 |
|
HW |
21.3 |
21.0 |
18.7 |
14.4 |
12.1 |
11.8 |
11.2 |
11.2 |
27.8 |
31.7 |
16.9 |
11.9 |
|
HL |
25.3 |
21.8 |
22.2 |
17.1 |
14.4 |
15.0 |
14.2 |
15.7 |
37.0 |
41.9 |
20.8 |
20.3 |
|
MN |
21.4 |
19.1 |
19.6 |
14.1 |
13.6 |
14.0 |
11.7 |
13.2 |
33.7 |
34.4 |
18.6 |
17.8 |
|
MFE |
18.6 |
15.8 |
15.8 |
10.3 |
9.2 |
11.3 |
10.7 |
9.8 |
26.8 |
29.4 |
15.3 |
14.1 |
|
MBE |
11.9 |
11.1 |
10.5 |
7.1 |
7.5 |
7.0 |
6.6 |
6.7 |
19.2 |
20.2 |
9.8 |
8.3 |
|
IFE |
7.4 |
6.2 |
6.4 |
6.3 |
5.6 |
5.5 |
5.2 |
6.1 |
13.7 |
14.8 |
8.0 |
8.4 |
|
IBE |
10.3 |
10.1 |
10.2 |
9.4 |
9.9 |
9.7 |
8.5 |
9.1 |
19.6 |
22.3 |
11.7 |
13.0 |
|
IN |
1.7 |
2.2 |
1.8 |
3.5 |
3.4 |
3.5 |
3.6 |
3.6 |
7.8 |
9.9 |
6.0 |
5.5 |
|
EN |
3.2 |
3.1 |
2.8 |
2.2 |
2.0 |
1.9 |
2.4 |
2.4 |
6.2 |
6.2 |
3.2 |
3.2 |
|
ED |
5.1 |
5.4 |
5.0 |
3.7 |
3.9 |
4.2 |
3.7 |
4.4 |
8.0 |
11.2 |
6.4 |
5.9 |
|
SN |
2.3 |
2.7 |
3.2 |
1.9 |
1.9 |
2.2 |
2.0 |
2.4 |
2.9 |
4.5 |
3.0 |
2.7 |
|
SL |
6.8 |
6.6 |
7.0 |
4.6 |
5.3 |
4.9 |
4.6 |
5.1 |
8.6 |
11.7 |
7.0 |
6.5 |
|
TYD |
4.8 |
4.9 |
3.8 |
2.4 |
2.1 |
2.4 |
1.9 |
2.3 |
4.8 |
5.4 |
3.1 |
2.4 |
|
TYE |
1.7 |
1.8 |
1.3 |
2.0 |
1.6 |
0.9 |
0.7 |
1.1 |
2.6 |
2.2 |
1.9 |
1.6 |
|
IUE |
2.2 |
2.1 |
1.8 |
3.7 |
3.1 |
2.8 |
2.6 |
4.6 |
4.1 |
7.9 |
4.4 |
4.3 |
|
UEW |
3.2 |
3.7 |
3.4 |
3.4 |
2.7 |
2.3 |
2.1 |
2.1 |
6.2 |
6.1 |
3.4 |
3.6 |
|
FLL |
8.9 |
8.6 |
9.4 |
9.4 |
7.9 |
5.3 |
5.2 |
6.1 |
16.0 |
19.7 |
9.2 |
10.1 |
|
HAL |
14.3 |
11.9 |
11.7 |
11.8 |
9.4 |
8.0 |
8.0 |
9.3 |
18.1 |
26.0 |
13.0 |
12.0 |
|
TFL |
10.4 |
7.2 |
7.9 |
7.2 |
6.6 |
4.5 |
4.3 |
6.3 |
10.5 |
17.9 |
7.7 |
8.0 |
|
FL |
30.6 |
27.4 |
24.7 |
16.3 |
17.0 |
15.2 |
14.8 |
16.9 |
37.1 |
40.1 |
20.8 |
21.0 |
|
TL |
26.7 |
24.6 |
20.4 |
17.8 |
16.0 |
18.4 |
16.4 |
19.0 |
42.9 |
49.7 |
24.4 |
25.6 |
|
TFOL |
40.0 |
33.9 |
32.6 |
27.3 |
24.0 |
23.8 |
21.9 |
26.1 |
57.6 |
59.1 |
34.3 |
34.4 |
|
FOL |
30.6 |
26.2 |
22.9 |
17.3 |
16.4 |
15.9 |
14.3 |
16.8 |
38.1 |
39.5 |
24.1 |
23.7 |
|
FTL |
24.4 |
22.2 |
21.3 |
13.2 |
14.5 |
12.5 |
10.1 |
13.4 |
33.0 |
32.1 |
20.5 |
19.8 |
SVL—Snout-vent length | HW—Head
width | HL—Head length | MN—Distance from the back of the mandible to the
nostril | MFE—Distance from the back of the mandible to the front of the eye |
MBE—Distance from the back of the mandible to the back of the eye |
IFE—Distance between the front of the eyes | IBE—Distance between the back of
the eyes | IN—Internasal space | EN—Distance from the front of the eye to the
nostril | ED—Eye diameter | SN—Distance from the nostril to the tip of the
snout | SL—Distance from the front of the eye to the tip of the snout |
TYD—Greatest tympanum diameter | TYE—Distance from tympanum to the back of the
eye | IUE—Minimum distance between upper eyelids | UEW—Maximum width of inter
upper eyelid | FLL—Forelimb length | HAL—Hand length | TFL—Third finger length
| FL—Femur length | TL—Tibia length | TFOL—Length of tarsus and foot | FOL—Foot
length | FTL—Fourth toe length.
Table 2. Samples used for
phylogenetic analysis with their GenBank accession numbers and other details.
|
Taxa |
Voucher number |
Locality |
Accession number |
Reference |
|
A. indoburmanensis |
MZMU 1650 |
Mizoram, India |
MT790757 |
- |
|
O. anlungensis |
HNNU 1008-109 |
China |
KF185049 |
Chen et al. 2013 |
|
O. yizhangensis |
SYSa001870 |
China |
KT315394 |
Wang et al. 2015 |
|
O. lungshengensis |
SYSa002229 |
China |
KT315395 |
Wang et al. 2015 |
|
O. lungshengensis |
806 |
China |
MH193554 |
Li et al. 2018 |
|
O. schmackeri |
HNNU 0908-349 |
China |
KF185047 |
Chen et al. 2013 |
|
O. fengkaiensis |
SYSa001025 |
China |
KT315382 |
Wang et al. 2015 |
|
O. hainanensis |
SYSa000636 |
China |
KT315384 |
Wang et al. 2015 |
|
O. wuchuanensis |
GZNU20180608019 |
China |
MW481358 |
Luo et al. 2021 |
|
O. jingdongensis |
SYSa002995 |
China |
KT315386 |
Wang et al. 2015 |
|
O. jingdongensis |
IEBR 3948 |
Vietnam |
KX893902 |
Ngo et al. 2016 |
|
O. margaretae |
SYSa002317 |
China |
KT315392 |
Wang et al. 2015 |
|
O. grahami |
CAS 207504 |
China |
DQ283241 |
Frost et al. 2006 |
|
O. mawphlangensis |
MZMU 2267 |
Mizoram, India |
MZ229896 |
- |
|
O. mawphlangensis |
MZMU 139 |
Mizoram, India |
MT814039 |
- |
|
O. mawphlangensis |
MZMU 138 |
Mizoram, India |
MT814038 |
- |
|
O. mawphlangensis |
MZMU 3021 |
Meghalaya, India |
OP979109 |
- |
|
O. mawphlangensis |
MZMU 3020 |
Meghalaya, India |
OP979112 |
- |
Table 3. The uncorrected K2p
-distance of 16S rRNA partial gene sequence among members of the Odorrana
sequences.
|
|
Taxon |
K2p distance |
||||||||||||||||
|
1 |
2 |
3 |
4 |
5 |
6 |
7 |
8 |
9 |
10 |
11 |
12 |
13 |
14 |
15 |
16 |
17 |
||
|
1 |
O. mawphlangensis_OP979109_Meghalaya |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
2 |
O. mawphlangensis_OP979112_Meghalaya |
0.004 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
3 |
O. mawphlangensis_MZ229896_Mizoram |
0.011 |
0.011 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
4 |
O. mawphlangensis_MT814039_Mizoram |
0.011 |
0.011 |
0.000 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
5 |
O. mawphlangensis_MT814038_Mizoram |
0.011 |
0.011 |
0.000 |
0.000 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
6 |
O. fengkaiensis_KT315382 |
0.110 |
0.110 |
0.096 |
0.096 |
0.096 |
|
|
|
|
|
|
|
|
|
|
|
|
|
7 |
O. hainanensis_KT315384 |
0.108 |
0.108 |
0.093 |
0.093 |
0.093 |
0.020 |
|
|
|
|
|
|
|
|
|
|
|
|
8 |
O. jingdongensis_KT315386 |
0.037 |
0.037 |
0.025 |
0.025 |
0.025 |
0.099 |
0.092 |
|
|
|
|
|
|
|
|
|
|
|
9 |
O. wuchuanensis_MW481358 |
0.049 |
0.049 |
0.036 |
0.036 |
0.036 |
0.091 |
0.083 |
0.034 |
|
|
|
|
|
|
|
|
|
|
10 |
O. margaretae_KT315392 |
0.051 |
0.051 |
0.039 |
0.039 |
0.039 |
0.120 |
0.112 |
0.022 |
0.043 |
|
|
|
|
|
|
|
|
|
11 |
O. yizhangensis_KT315394 |
0.083 |
0.083 |
0.070 |
0.070 |
0.070 |
0.112 |
0.099 |
0.068 |
0.060 |
0.073 |
|
|
|
|
|
|
|
|
12 |
O. lungshengensis_KT315395 |
0.078 |
0.078 |
0.065 |
0.065 |
0.065 |
0.088 |
0.076 |
0.063 |
0.055 |
0.073 |
0.041 |
|
|
|
|
|
|
|
13 |
O. grahami_DQ283241 |
0.042 |
0.042 |
0.029 |
0.029 |
0.029 |
0.110 |
0.102 |
0.018 |
0.034 |
0.022 |
0.078 |
0.073 |
|
|
|
|
|
|
14 |
O. schmackeri_KF185047 |
0.093 |
0.093 |
0.079 |
0.079 |
0.079 |
0.076 |
0.073 |
0.092 |
0.084 |
0.102 |
0.104 |
0.086 |
0.087 |
|
|
|
|
|
15 |
O. jingdongensis_KX893902 |
0.039 |
0.039 |
0.027 |
0.027 |
0.027 |
0.102 |
0.094 |
0.002 |
0.036 |
0.020 |
0.070 |
0.065 |
0.020 |
0.095 |
|
|
|
|
16 |
O. lungshengensis_MH193554 |
0.078 |
0.078 |
0.065 |
0.065 |
0.065 |
0.088 |
0.076 |
0.063 |
0.055 |
0.073 |
0.041 |
0.000 |
0.073 |
0.086 |
0.065 |
|
|
|
17 |
O. anlungensis_KF185049 |
0.082 |
0.082 |
0.068 |
0.068 |
0.068 |
0.105 |
0.100 |
0.071 |
0.068 |
0.079 |
0.061 |
0.056 |
0.076 |
0.100 |
0.069 |
0.056 |
|
|
18 |
Amolops indoburmanensis_MT790757 |
0.186 |
0.186 |
0.168 |
0.168 |
0.168 |
0.180 |
0.177 |
0.171 |
0.165 |
0.177 |
0.160 |
0.161 |
0.172 |
0.186 |
0.168 |
0.161 |
0.171 |
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